A central current debate in community ecology concerns the relative importance of deterministic versus stochastic processes underlying community structure. However, the concept of stochasticity presents several profound philosophical, theoretical and empirical challenges, which we address here. The philosophical argument that nothing in nature is truly stochastic can be met with the following operational concept of neutral stochasticity in community ecology: change in the composition of a community (i.e. community dynamics) is neutrally stochastic to the degree that individual demographic events -birth, death, immigration, emigration -which cause such changes occur at random with respect to species identities. Empirical methods for identifying the stochastic component of community dynamics or structure include null models and multivariate statistics on observational species-by-site data (with or without environmental or trait data), and experimental manipulations of 'stochastic' species colonization order or relative densities and frequencies of competing species. We identify the fundamental limitations of each method with respect to its ability to allow inferences about stochastic community processes. Critical future needs include greater precision in articulating the link between results and ecological inferences, a comprehensive theoretical assessment of the interpretation of statistical analyses of observational data, and experiments focusing on community size and on natural variation in species colonization order.

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Resource pulses within structured communities can lead to changes in the ecological roles of community members, particularly for species that exhibit plasticity in resource use. The red‐breasted nuthatch Sitta canadensis is a facultative excavating cavity‐nester that forages on seeds and insects, thus exhibits plasticity in both nesting habits and diet. In a long‐term study of cavity‐nesting vertebrates, we used point counts, and nest and vegetation surveys to examine the effects of two resource pulses of mountain pine bark beetle prey Dendroctonus ponderosae and tree cavities via excavator populations, on population densities and cavity reuse of red‐breasted nuthatches, from 1997–2006. We observed a doubling in mean nut‐hatch densities from 0.12 to 0.24 individuals ha−1 then a collapse later in the decade to 50% below endemic levels (0.06 individuals ha−1). These regional fluctuations were positively correlated with densities of trees recently infected by bark beetles. Because nuthatches range over large areas in winter, this regional correlation suggests that populations responded to the increase in winter food supply. At the site scale, general linear mixed effects models showed that nuthatch populations increased following years of high densities of downy woodpeckers Picoides pubescens, suggesting that downy woodpeckers were important facilitators, via cavity excavation. Type of nesting cavity used by nuthatches varied during the pulse, such that the use of existing cavities (in lieu of excavation) increased on sites that had harboured downy woodpecker nests in the previous year. We conclude that increased densities of cavity excavators allowed facultative excavators to increase their reuse of cavities, which may have contributed to the dramatic increases in nuthatch populations. However, nuthatch populations collapsed after the boom suggesting that this dual resource pulse may have destabilized populations by enabling densities to reach unsustainable levels. Thus, plasticity in resource use can have serious costs as well as benefits.

Secondary cavity-nesting birds depend on tree cavities for nesting and roosting, but many studies of these birds are conducted using nest boxes. Implementation of effective conservation strategies for cavity-nesting species such as nest-site supplementation requires careful comparisons of fecundity and other vital rates for birds using both natural and artificial nest site types. We compared breeding phenology, clutch and brood sizes, and fledging success of Tree Swallows (Tachycineta bicolor) nesting in tree cavities and nest boxes during 2001–2003 in British Columbia, Canada. Swallows using nest boxes initiated egg-laying and hatched young at approximately the same time as those in tree cavities (2 June, 23 June, respectively). Female Tree Swallows in boxes laid larger clutches (5.9 ± 0.9 eggs, N = 76) than those in tree cavities (4.2 ± 1.6 eggs, N = 67). The mean number of nestlings hatched was greater in nest boxes (5.2 ± 1.1 nestlings, N = 67) than in tree cavities (2.6 ± 2.0 nestlings, N = 58). Pairs in boxes were over twice as successful in producing fledglings (93.4%; 57 of 61 pairs fledged > 1 young) than those in tree cavities (35.8%; 19 of 53 pairs). Of those successful nests, pairs nesting in boxes fledged 5.1 ± 1.1 young (N = 57), whereas those in tree cavities fledged 3.5 ± 1.2 young (N = 18). Because cavities in nest boxes averaged 60% larger in volume and 1.8 cm wider internally than tree cavities, we suggest that increased reproductive output was correlated with boxes enabling a larger clutch size. In previous research, we found that Tree Swallows were a poor competitor with other cavity-nesting passerines for tree cavities. The addition of nest boxes may serve as an effective way to supplement local reproduction for secondary cavity-nesting bird populations by reducing competition for limited nest sites. This is especially true in regions where the availability of natural nesting sites is highly variable, and where species compete with many other cavity-nesting passerines using a similar ecological niche and nesting cavities.

Woodpeckers and other excavators create most of the holes used by secondary cavity nesters (SCNs) in North American temperate mixedwood forests, but the degree to which excavators release SCNs from nest‐site limitation is debated. Our goal was to quantify how excavators maintain the diversity and abundance of secondary cavity nesters in a temperate forest through the creation of tree cavities. We examined the short‐ and long‐term (legacy) effects of excavators (principally woodpeckers, but also red‐breasted nuthatches and black‐capped chickadees) on forest biodiversity using longitudinal monitoring data (1,732 nest cavities, 25 sites, 16 years) in British Columbia, Canada. Sites with higher densities of excavator nests had more cavities available, higher species richness of SCNs and higher nest density of SCNs, indicating the importance of a standing stock of cavities. Years with higher nesting densities of excavators were followed by years with higher SCN diversity, indicating that the creation of nesting opportunities through fresh excavation releases SCNs from community‐wide nest‐site limitation. We also show that excavators leave a ‘legacy’ of biodiversity (species richness and abundance) at a site by accumulating cavities at rates faster than they become unusable by decay or destruction. By quantifying site‐level effects of cavity excavation on the SCN community, our study highlights the key role of excavators as ecosystem engineers that maintain forest wildlife biodiversity.